J. South Asian Nat. Hist., ISSN 1022-0828. May, 1999. Vol.4, No. 1, pp. 71-81, 6 figs., 1 tab. © Wildlife Heritage Trust of Sri Lanka, 95 Cotta Road, Colombo 8, Sri Lanka.
Notes on the taxonomy of two species of swimming crabs, Charybdis smithii MacLeay, 1838, and C. omanensis Leene, 1938 (Crustacea: Decapoda: Brachyura: Portunidae) from the Indian Ocean
Peter K. L. Ng * & Masatsune Takeda **
* Department of Biological Sciences, National University of Singapore, Kent Ridge, Singapore 119260, Republic of Singapore. ** Department of Zoology, National Science Museum, 3-23-1 Hyakunin- cho, Shinjuku-ku, Tokyo 169/ Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Tokyo 113, Japan.
Abstract Two supposedly closely allied species of swiming crabs, Charybdis smithii MacLeay, 1838, and C. omanensis Leene, 1938, are compared. The two species differ in many carapace, chelipedal, abdominal, thoracic sternal and gonopodal characters. Swarming in Charybdis smithii is again reported, and feeding on krill is suggested as the reason for this unusual behaviour.
Introduction Collections by the Japanese research vessel Family Portunidae WAKADORI-MARU off the Arabian Sea recently ob tained specimens of a swimming crab which was Charybdis omanensis Leene, 1938 swarming near the surface. A study of these speci (Figures 1,2) mens showed that they belong to Charybdis smithii MacLeay, 1838. As there has been some discussion ? Charybdis (Goniohellenus) hoplites, Rathbun, 1911: 207 over the possible synonymy of this species with C. Charybdis (Goniohellenus) hoplites var. omanensis Leene, 1938: 104, Fig. 57. omanensis Leene, 1938, and we had an opportunity to Charybdis (G.) hoplites omanensis, Stephenson, 1972: 35. compare specimens of both species, the present note Charybdis (Goniohellenus) hoplites omanensis, Zarenkov, makes additional comments on their supposed affini 1968: 34, fig. 1, 2; Zarenkov, 1971: 181. ties and possible synonymy. Charybdis (Goniohellenus) omanensis, Tiirkay, 1986: 156, The terms used here essentially follow those by Figs. 43-45. Wee & Ng (1995). The abbreviations G1 and G2 are used for the male first and second pleopods respec Material. 1 male (cw 37.3 mm, cl 27.3 mm), 4 females tively Measurements are of the carapace width (cw) (largest cw 42.9 mm, cl 28.3 mm) (USNM 234212); and length (cl). Specimens examined are deposited in Indian Ocean, Gulf of Aden, Somalia, Haso, the National Science Museum, Tokyo (NSMT); Na 11Q42'36"N 50Q21'24,/E, trawl, depth 180 m; coll. R/ tional Museum of Natural History (USNM), V BEINTA, 16 V I1987. Smithsonian Institution, Washington D.C.; Zoologi cal Reference Collection (ZRC), National University Discussion. In the USNM are several specimens of C. of Singapore; MacLeay Museum, University of Syd omanensis collected from the Gulf of Aden. They agree ney (MMUS); and the Nationaal Naturhistorisch very well with the detailed descriptions and figures Museum (ex Rijksmuseum van Natuurlijke Historie, of the species provided by Leene (1938). The only RMNH). major difference we could discern was that in the N g & T akeda
Figure 1. Charybdis omanensis. Male (cw 37.3 view; b, ventral view; c, outer surface of mm, 27.3 mm) (USNM 234212). a, Dorsal chela.
J. South Asian Nat. Hist. Charybdis smithii & C. omanensis
Figure 2. Charybdis omanensis. Male (cw 37.3 mm, 27.3 antennal segment; c, abdominal segments 3-6 and telson; mm) (USNM 234212). a, left third maxilliped; b, right basal d, left G l; e, left G2. present specimens, the median frontal teeth are all Goniosoma truncata, A. Milne Edwards, 1861: 380, pi. 34 fig. sharp and not blunt. Additional figures of several 4 (not Fabricius, 1798). useful characters diagnosing C. omanensis are pro Charybdis (Goniohellenus) edwardsi Leene & Buitendijk, vided here (Figs. 1, 2) to supplement the otherwise 1949: 296, Figs. 3, 4c; Sankarankutty & Rangarajan, excellent description and figures of Leene (1938). 1964:311; Della Croce & Holthuis, 1965:33; Losse, 1969: 145; Silas, 1969: 26, Fig. 11, PI. 3 figs. A, B; Heath, 1971: Charybdis omanensis has been reported from vari 1; Daniel & Chakrapany, 1984: 101. ous parts of the northern Indian Ocean, viz. the Gulf Charybdis edwardsi, Stephenson, 1967: 9. of Oman (Leene, 1938), Gulf of Aden (Zarenkov, 1968, Charybdis (Goniohellenus) smithii, Stephenson, 1972: 35. 1971), Red Sea (Zarenkov, 1971; Tiirkay, 1986) and Charybdis (Goniohellenus) smithii, Crosnier & Thomassin, Gulf of Bengal (Zarenkov, 1968). 1974: 1109, Fig. 7. Charybdis (Goniohellenus) smithi - Spiridonov, 1994: 133. Charybdis smithii MacLeay, 1838 Charybdis smithi, Zamorov et al., 1991a: 39; Zamorov et (Figures 4-6) al., 1991b: 70.
Charybdis smithii MacLeay, 1838: 61; Krauss, 1843: 24; Material. Holotype female (cw 56.4 mm, cl 40.1 mm) Stebing, 1910: 37; Stephenson & Rees, 1967a: 9; (dried) (MMUS), MacLeay Collection. 1 male, 1 female Stephenson & Rees, 1967b: 285; Griffin & Stanbury, (NSMT), 2 males (cw 44.1 mm, cl 31.9 mm, cw 58.3 1970: 123; Van Couwelaar et al., 1997: 1251. mm, cl 41.2 mm), 1 female (cw 52.6 mm, cl 36.4 mm)
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(ZRC); station 5, larval nets targetting squids, net 2 m stronger spines; posterior margin finely granulated; diameter, surface trawl, 15 minutes trawl time, Ara 2 anterior surfaces gently rugose to smooth. Carpus bian Sea, 16Q00'N 64Q00'E; coll. WAKADORI-MARU, with granulated ridge ending in strong inner spine; 14 X 1996.1 male, 1 female (NSMT); station 3, larval outer angle with 3 small but distinct spines, posterior nets targetting squids, net 2 m diameter, surface trawl, 2 transversely in line with each other. Manus with 4 15 minutes trawl time, Arabian Sea, 14Q00'N 64Q00'E; spines; 1 strong, slightly hooked spine at carpal- coll. WAKADORI-MARU, 12 X 1996. (Specimens ex manus articulation; inner margin of upper surface amined which have been previously reported by with 1 spine; upper margin with 2 distal spines just Stephenson (1967) and Stephenson & Rees (1967a) before movable finger; manus with 7 longitudinal from the USNM are not listed again here) granular costae, 6 on outer face and margins, and 1 on inner face; outer surface of each finger with 2 Description. Carapace evenly convex longitudinally smooth, longitudinal costae. Cutting margins of fin and transversely, surface smooth; very short pubes gers with numerous teeth and denticles. Movable fin cence present on anterolateral and frontal regions, gers shorter than manus. and along carapace grooves and depressions. Only First to third ambulatory legs similarly shaped; epibranchial crista visible, being distinct along proxi posterior margins of meri lined with very small, mal half and becoming very weak to undiscernible rounded granules, distal angle of posterior margin along distal part; all other ridges absent. Front cut with sharp spine; propodus with submedian longi into 6 acutely triangular teeth, lateral teeth largest, tudinal sulcus, distal margins with numerous small, margins of all teeth lined with small rounded gran sharp granules; dactylus substyliform, with 2 deep ules; median 2 teeth anterior of rest of frontal teeth, longitudinal grooves. Fourth leg natatory; dactylus separated by distinct cleft; median teeth separated with weak median carina, tip with sharp tooth; pos from submedian teeth by distinct, shallow, broad U- terior margin of propodus with about 5 small spines shaped cleft; submedian teeth sharp, separated from or sharp granules, anterior surface with longitudinal lateral teeth by deep V-shaped cleft. Supraorbital mar groove; merus short, with faint median longitudinal gin finely granulated, divided into 3 lobes by 2 dis ridge, strong posterodistal spine anterior to distal tinct, median notches. Anterolateral margin arcuate, spine on posterior margin. with 5 teeth, all separated by distinct deep, U-shaped Surfaces of thoracic sternites slightly rugose to clefts; first to fourth teeth truncate, second tooth small smooth, glabrous; suture between sternites 4 and 5, est, margins of these teeth lined with distinct rounded and sternites 2 and 3 gently granulose. Male abdo and sharp granules; fifth tooth triangular, anterior men triangular; third to fifth segments completely margin lined with small, sharp granules, posterior fused without trace of sutures, lateral margin of third margin smooth, with epibranchial crest starting me and fourth segments separated by deep V-shaped cleft, dially. Posterolateral margins smoth, gently concave, fourth segment with low, median transverse ridge; converging towards posterior margin of carapace; sixth segment trapezoidal, lateral margins gently sinu posterior carapace margin forming distinct angular ous to slightly convex outwards; telson triangular, junction with end of posterolateral margin. Posterior lateral margins gently convex outwards. carapace margin entire, gently convex posteriorly. Gl curving outwards; proximal part gently taper Orbits relatively large; eyes large, cornea well devel ing towards bending point; distal half gently sinu oped, filling most of orbit. Infraorbital lobe large, dis ous, tip gently curved downwards; basal lobe trian tinctly visible from dorsal view. Suborbital, sub- gular. G2 with very short distal segment. branchial and subhepatic regions finely rugose, cov ered with short pubescence. Basal antennal segment Discussion. See below for detailed discussion. slightly wider than orbit, granulated throughout, with Charybdis smithii has been reported from a wide area 2 transverse rows of large, rounded granules, one near in the Indian Ocean (see Van Couwelaar et al., 1997). the antenna, the other adjacent to the base. Ischium of third maxilliped rectangular, with deep General Discussion oblique sulcus; anterointernal angle granulated; Various authors had commented that Charybdis merus triangular, anteroexternal angle strongly smithii MacLeay, 1838, was affiliated or synonymous auriculiform, reaching above tip of exopod, inner an with Charybdis (Goniohellenus) hoplites var. omanensis gle granulated; exopod broad, about half width of is Leene, 1938, and/or Charybdis (Goniohellenus) edwardsi chium, inner margin finely granulated. Leene & Buitendijk, 1949 (Stephenson, 1967a, b, 1970; Chelipeds subequal in size, between 2-3 times Stephenson & Rees, 1967a). All three species had been length of carapace. Outer anterior surface of merus described from single specimens. Charybdis smithii rugose; anterior margin finely denticulated, with 2 was described from a single female from the Cape of
74 J. South Asian Nat. Hist. Charybdis smithii & C. omanensis
Figure 3. Charybdis smithii. Holotype male view; b, carapace; c, frontal margin, (cw 56.4 mm, cl 40.1 mm) (MMUS). a, overall
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Figure 4. Charybdis smithii. Holotype male (cw 56.4 last three ambulatory legs; c, dorsal margin of chela; mm, cl 40.1 mm) (MMUS). a, third maxillipeds; b, d, outer surface of chela
J. South Asian Nat. Hist. Charybdis smithii & C. omanensis
Figure 5. Charybdis smithii. Male (cw 44.1 mm, cl outer surface of chela. 31.9 mm) (ZRC). a, Dorsal view; b, ventral view; c,
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Figure 6. Charybdis smithii. Male (cw 44.1 mm, cl 31.9 mm) leg; e, right fourth ambulatory leg; /, left Gl; g, left G2; h, (ZRC). a, right side of carapace; b, left third maxilliped; c, abdominal segments 3-6 and telson. right basal antennal segment; d, right third ambulatory
78 J. South Asian Nat. Hist. Charybdis smithii & C. omanensis
Table 1. Differences between Charybdis omanensis Leene, 1938, and Charybdis smithii MacLeay, 1838.
Charybdis omanensis Charybdis smithii
carapace most of dorsal surface covered only anterolateral and frontal regions, with pubescence as well as depressed areas and groove covered with fine pubescence
epibranchial region sharp and distinct throughout length, relatively sharp along anterior half, lined with small, rounded granules very weak to almost undiscernible along posterior half, smooth along entire length
protogastric, mesogastric with distinct and transverse cristae smooth, with no cristae or granules metagastric regions which are lined with small, rounded granules
mesobranchial and with numerous small,rounded granules; smooth; mesobranchial regions gently cardiac regions mesobranchial regions distinctly inflated convexbut not highly inflated
posterior margin of almost straight from dorsal view, sinuous gently convex outwards from dorsal carapace from posterior view view, straight from posterior view
fifth anterolateral tooth acutely triangular, distinctly longer relatively small, subequal in length than fourth anterolateral tooth to fourth anterolateral tooth
costae on chelipedal each costa composed of 1-3 tightly each costa composed of only 1 distinct manus packed rows small or flattened of row of granules; areas between costae granules; areas between costae gently rugose distinctly rugose
spines on chelipeds strong weak
posterior face of granulose to strongly rugose slightly rugose to smooth merus of cheliped
surface of thoracic rugose to granulose, pubescent gently rugose (anterior sternites) sternites to smooth (posterior sternites), glabrous
suture between distinctly granulose slightly granulose sternites 3 and 4
suture between distinctly granulose slightly granulose sternites 4 and 5
male abdominal second, third and fourth segments with second, third and fourth segments with segments strong, transverse median keel (that on low ridge which is not cristate or fourth segment laterally interrupted) keeled
ambulatory legs marginal and/or median costae on marginal and/or median costae on dactylus, propodus and merus strong, dactylus, propodus and merusrelatively broad; areas between these costae weaker, narrow; areas between these pubescent costae glabrous
Gl strongly curving gently curving outwards outwards
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Good Hope (South Africa), C. (G.) hoplites var. ters. Some of the specimens, however, were subadults, omanensis from a single male specimen from the Gulf suggesting that this particular swarming might not of Oman, and C. (G.) edwardsi from a single dried male have been associated with breeding. The mouthparts from Malabar, east India. The problems arose because of almost all the specimens examined had clumps of MacLeay's description was inadequate and the sub krill in various stages of digestion. It appears that as sequent description of the species by Barnard (1950) the freshly collected specimens were killed and pre was based primarily on observations of the type made served, they regurgitated thair last meal, which co for him by M. Ward. Finally, Stephenson & Rees agulated around their mouthparts. It therefore sug (1967b) argued that Charybdis smithii MacLeay, 1838, gests that the present crabs were swarming to feed on was synonymous with Charybdis (Goniohellenus) swarms of krill near the ocean surface. As far as pub edwardsi Leene & Buitendijk, 1949. lished records go, C. omanensis does not show this The present specimens of C. smithii on hand agree surface swarming behaviour. well with the descriptions of previous authors. Barnard (1950) and Stephenson & Rees (1967c) had Acknowledgements commented that there was weak granulation on the The authors are most grateful to Akihiko Yatsu of the carapace of C. smithii. In the series of specimens on National Research Institute of Far Seas Fisheries who hand, however, there are no visible granules anywhere kindly collected the recent specimens and passed on the carapace. Some parts of the metabranchial re them to us for this study. Thanks are due to Ray Man gions may appear somewhat rough and the anterior ning and Rafael Lemaitre (USNM), and Lipke part of the epibranchial cristae may feel uneven, but Holthuis and Charles Fransen (RMNH) for their ex no granules can be discerned. Crosnier & Thomassin cellent hospitality during the first author's visits to (1974) provided excellent figures of this species, their institutes. Shane Ahyong (Australian Museum) which the present specimens agree with excellently kindly examined and photographed the holotype of The holotype is refigured here. Although it is in poor Charybdis smithii for the first author. Vasily Spiridonov condition, most of the diagnostic characters for the kindly provided several interesting papers on the bi species identified here are still clearly discernible ology of C. smithii. Lipke Holthuis kindly reviewed (Figs. 4,5) the manuscript. The study has been partially sup Leene (1938) separated C. hoplites omanensis from ported by a 1997 travel grant to the first author from C. hoplites hoplites using a number of carapace and the National University of Singapore. cheliped features. She also referred specimens re ported by Rathbun (1911) as "? Charybdis Literature cited (Goniohellenus) hoplites" from Saya de Malha in the Indian Ocean as belonging to her species. Stephenson Barnard, K. H. 1950. Descriptive catalogue of South African (1967: 10) had suggested that C. hoplites omanensis Decapod Crustacea. Ann. S. Afr. Mus., 38: 1-837. might be a synonym of C. smithii and casted doubt on Crosnier, A. & B. Thomassin. 1974. Sur des crabes de la the validity of C. hoplites omanensis in Stephenson & famille Portunidae (Crustacea Decapoda) nouveaux pour Madagascar ou rares. Bull. Mus. Natn. Hist. Nat., Rees (1967:286) and Stephenson (1972:35). Zarenkov Paris, (3)165(241): 1097-1118. (1968,1971), however, continued to use the name C. Daniel, A. & S. Chakrapany, 1984. Observations on the hoplites omanensis, with Tiirkay (1986) finally recog swarming, breeding habits and some larval stages of nising it as a distinct species. We concur with the deep-sea portunid crab Charybdis (Goniohellenus) Turkay's (1968) decision. After examining the series edwardsi Leene et Buitendijk, 1949 in the northern of specimens of both C. smithii and C. omanensis, in Arabian Sea and off the Madras coast during January cluding small ones and of both sexes, we have noted to March 1976 - to 1979. Rec. Zool. Surv. India, 12:101- many differences which clearly support their separa 108. tion as two distinct species (Table 1). Most are in ad Della Croce, N. & L. B. Holthuis. 1965. Swarming of Charybdis (Goniohellenus) edwardsi Leene & Buitendijk dition to the often cited difference of whether the ma in the Indian Ocean (Crustacea Decapoda, Portunidae). jor regions or cristae on the dorsal surface of the cara Boll. Mus. Inst. Biol. Univ. Genova, 33(199): 33-38. pace are granulated or smooth. Fabricius, J. C. 1798. Entomologia Systematica Emendata It is well known that C. smithii swarms near the et Aucta Secundum Classes, Ordines, Genera, Species, surface and Van Couwelaar et al. (1997) provides an Adjectis Synonymis, Locis, Observationibus, excellent review and study of this phenomenon. The Discriptionibus. Supplementum Entomologiae present specimens were also caught while they were Systematicae. pp. 572. Hafniae. swarming just below the surface. There was no land Griffin, D. J. G. & P. J. Stanbury. 1970. Type specimens in nearby, and the crabs had either risen from deep wa the MacLeay Museum, University of Sydney. V. Decapod Crustaceans. Proc. Linn. Soc. N. S. W., 95(1): ter or had migrated some distance from shallow wa 122-131.
80 J. South Asian Nat. Hist. Charybdis smithii & C. omanensis
Heath, J. R. 1971. Crabs of Dar es Salaam: Part 1, Stebbing, T. R. R. 1910. General catalogue of South African Portunidae. Tanzania Notes Rec., 72:1-17. Crustacea, (part V of S. A. Crustacea, for the Marine Krauss, R 1843. Die Stidafrikanischen Crustaceen. Eine Investigations in South Africa). Ann. S. Afr. Mus., 6: Zusammenstellung aller bekanten malacostraca, 281-599,8 pis. Bemerkungen iiber deren Lebensweise und Stephenson, W. 1967. The portunid crabs (Crustacea: geographische Verbreitung, nebst. Beschreibung und Portunidae) collected by the NAGA expedition. Naga Abbildung mehrer neuen arten. 68 pp, 4 pis. Rep., 4: 4-37, 4 pis. Leene, J. E. 1938. The Decapod Brachyura of the Siboga Stephenson, W. 1972. An annotated check list and key to Expedition. VII. Brachygnatha: Portunidae. Monogr. the Indo-West Pacific swimming crabs (Crustacea: Siboga, 39C3(131): 1-156. Decapoda: Portunidae). Roy. Soc. N. Z. Bull., 10:1-64. Leene, J. E. & A. M. Buitendijk. 1949. Notes on Charybdis Stephenson, W. & M. Rees. 1967a. Portunid crabs from ihlei nov. spec., Charybdis beauforti nov. spec., and the International Indian Ocean Expedition in the Charybdis edwardsi nov. spec., from the collections of Smithsonian Collections (Crustacea: Portunidae). Proc. the British Museum (Natural History), London. Bijd. U. S. Natn. Mus., 122(3599): 1-33. Dierk., 28: 291-298. Stephenson, W. & M. Rees. 1967b. Portunid crabs collected Losse, G. R 1969. Notes on the portunid crab Charybdis by the 'Discovery' in the Indian Ocean. J. nat. Hist., 1: edwardsi Leene & Buitendijk, 1949, from the Western 285-288. Indian Ocean. J. nat. Hist., 3:145-152,1 pi. Tiirkay, M. 1986. Crustacea Decapoda Reptantia der Tiefsee MacLeay, W. S. 1838. On the Brachyurous Decapod des Roten Meeres. Senckenbergiana marit., 18: 123- Crustacea. Brought from the Cape by Dr. Smith, in A. 185. Smith, Illustrations of the Zoology of South Africa; Van Couwelaar, M., M. V. Angel & L. P. Madin. 1997. The consisting chiefly of figures and descriptions of the distribution and biology of the swimming crab objects of natural history collected during an Charybdis smithii McLeay, 1838 (Crustacea; Brachyura; expedition into the interior of South Africa, in the years Portunidae) in the NW Indian Ocean. Deep Sea Res., 1834,1835, and 1836; fitted out by 'The Cape of Good Pt II, Trop. Stud. Oceanogr., 44(6-7): 1251-1280. Hope Association for Exploring Central Africa:' Wee, D. P. C. & P. K. L. Ng. 1995. Swimming crabs of the together with a summary of African Zoology, and an genera Charybdis De Haan, 1833, and Thalamita inquiry into the geographical ranges of species in that Latreille, 1829 (Crustacea: Decapoda: Brachyura: quarter of the glode. Published under the Authority Portunidae) from Peninsular Malaysia and Singapore. of the Lords Commissioners of Her Majesty's Raffles Bull. Zool., Suppl. 1,128 pp. Treasury, Invertebratae (London: Smith, Elder and Zamorov, V. V., V. A. Spiridonov & G. P. Rudnev. 1991a. Co.), [1849], pp. 53-71, pis 2, 3. On the mass migration to the pelagial and the Milne Edwards, A. 1861. Etudes zoologiques sur les distribution of the Indian Ocean swimming crab Crustac,s r,cents de la famille des Portuniens. Arch. Charybdis smithi McLeay (Crustacea, Portunidae) Mus. Hist, nat., Paris, 10: 309-428. during the pelagic phase of its life-cycle. Zool. Zh., 70: Rathbun, M. J. 1911. Marine Brachyura. The Percy Sladen 39-42. Trust Expedition to the Indian Ocean in 1905, under Zamorov, V. V., V. A. Spiridonov & G. V. Napadovsky, the Leadership of Mr J. Stanley Gardiner, volume 3, 1991b. On the role of swimming crab Charybdis smithi number 11. Trans. Linn. Soc. London, (2) Zool., 14(2): McLeay in the feeding habit of yellowfin tuna Thunnus 191-261, pis. 15-20. albacares (Bonnaterre). Collective volume of working Sankarankutty, C. & K. Rangarajan. 1964. On a record of documents. Workshop on stock assessment of Charybdis (Goniohellenus) edwardsi Leene and yellowfin tuna in the Indian Ocean, FAO-UNDP, Rome, Buitendijk. J. mar. Biol. Ass. India, 6(2): 311. 6: 70-75. Silas, E. G. 1969. Exploratory fishing by the R. V. Varuna. Zarenkov, N. A. 1968. Crabs of Charybdis de Haan from Bull. Cent. Mar. Fish. Res. Inst., 12:1-86, pis. 1-15. collections of Soviet Expedition in the Pacific and Indian Spiridonov, V. A., 1994. The swimming crabs (Crustacea, Oceans. Vest, moskovs. Univ., Biol.-Zool., (6)2: 32-38. Brachyura, Portunidae) of submerged rises and insular Zarenkov, N. A. 1971. On the species-composition and shelves of the Atlantic and Indian Oceans. In: Bottom ecology of the Red sea decapod Crustacea. In: Benthos Fauna of Seamounts. Trans. P. P. Shirshov Inst. of the Shelf of the Red Sea, pp. 155- 203. Oceanol., Moscow, 129: 126-151.
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